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131-596: In 2013, researchers confirmed earlier speculation that the cause of the plague of Justinian was Yersinia pestis , the same bacterium responsible for the Black Death (1346–1353). Ancient and modern Yersinia pestis strains are closely related to the ancestor of the Justinian plague strain that has been found in the Tian Shan , a system of mountain ranges on the borders of Kyrgyzstan , Kazakhstan , and China , suggesting that

262-612: A GTPase-activating protein for members of the Rho family of GTPases such as RAC1 . YopT is a cysteine protease that inhibits RhoA by removing the isoprenyl group , which is important for localizing the protein to the cell membrane . YopE and YopT has been proposed to function to limit YopB/D-induced cytolysis. This might limit the function of YopB/D to create the pores used for Yop insertion into host cells and prevent YopB/D-induced rupture of host cells and release of cell contents that would attract and stimulate immune system responses. YopJ

393-608: A Lombard Kingdom in north and central Italy, which reached its zenith under the eighth-century ruler Liutprand . In 774, the kingdom was conquered by the Frankish king Charlemagne and integrated into the Frankish Empire . However, Lombard nobles continued to rule southern parts of the Italian peninsula well into the eleventh century, when they were conquered by the Normans and added to

524-575: A betrayal, Thurisind made an alliance with the Kutrigurs who devastated Moesia before end of the armistice. The Langobard and Roman army joined together and defeated the Gepids in 551. In the battle, Audoin 's son, Alboin killed Thurisind 's son, Turismod . In 552, the Byzantines, aided by a large contingent of Foederati , notably Lombards, Heruls and Bulgars, defeated the last Ostrogoths led by Teia in

655-499: A champion of freedom. In 47, a struggle ensued amongst the Cherusci and they expelled their new leader, the nephew of Arminius, from their country. The Lombards appeared on the scene with sufficient power to control the destiny of the tribe that had been the leader in the struggle for independence thirty-eight years earlier, for they restored the deposed leader to sovereignty. To the south, in 166 Cassius Dio reported that just before

786-513: A clinical isolate in the United States). In 2006 the genome sequence of a strain of biovar Antiqua was completed. Some strains are non-pathogenic, such as that of strain 91001 , whose sequence was published in 2004. Like Y. pseudotuberculosis and Y. enterocolitica , Y. pestis is host to the plasmid pCD1. It also hosts two other plasmids, pPCP1 (also called pPla or pPst) and pMT1 (also called pFra) that are not carried by

917-685: A cult of Odin, thus creating a conscious tribal tradition. Fröhlich inverts the order of events in Priester and states that with the Odin cult, the Lombards grew their beards in resemblance of the Odin of tradition and their new name reflected this. Bruckner remarks that the name of the Lombards stands in close relation to the worship of Odin, whose many names include "the Long-bearded" or "the Grey-bearded", and that

1048-564: A detrimental effect on non-human mammals; in the United States, these include the black-tailed prairie dog and the endangered black-footed ferret . Y. pestis is a non-motile coccobacillus , a facultative anaerobic bacterium with bipolar staining (giving it a safety pin appearance) that produces an antiphagocytic slime layer. Similar to other Yersinia species, it tests negative for urease , lactose fermentation, and indole . The species grows best in temperatures of 28–30 °C, and at

1179-511: A disease of rats; villagers in China and India asserted that when large numbers of rats were found dead, plague outbreaks soon followed. In 1898, French scientist Paul-Louis Simond (who had also come to China to battle the Third Pandemic) discovered the rat–flea vector that drives the disease. He had noted that persons who became ill did not have to be in close contact with each other to acquire

1310-485: A hospital in Beijing's Chaoyang district, prompting fears of an outbreak. The patient was a middle-aged man with fever, who had complained of difficulty breathing for some ten days, accompanied by his wife with similar symptoms. Police quarantined the emergency room at the hospital and controls were placed on Chinese news aggregators. On 18 November a third case was reported, in a 55-year-old man from Xilingol League , one of

1441-454: A mammalian host. While in the insect vector, proteins encoded by hemin storage system genetic loci induce biofilm formation in the proventriculus , a valve connecting the midgut to the esophagus . The presence of this biofilm seems likely to be required for stable infection of the flea. Aggregation in the biofilm inhibits feeding, as a mass of clotted blood and bacteria forms (referred to as "Bacot's block" after entomologist A.W. Bacot ,

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1572-400: A massive effect on prairie food webs. However, the transmission dynamics within prairie dogs do not follow the dynamics of blocked fleas; carcasses, unblocked fleas, or another vector could possibly be important, instead. The CO92 strain was isolated from a patient who died from pneumonia and who contracted the infection from an infected cat. In other regions of the world, the reservoir of

1703-458: A mild diarrhea, that reclassification would generate confusion. A comprehensive and comparative proteomics analysis of Y. pestis strain KIM was performed in 2006. The analysis focused on growth under four different sets of conditions that were designed to model flea and mammal hosts. Numerous bacterial small noncoding RNAs have been identified to play regulatory functions. Some can regulate

1834-450: A now classic experiment, Simond demonstrated how a healthy rat died of the plague after infected fleas had jumped to it from a rat that had recently died of the plague. The outbreak spread to Chinatown, San Francisco, from 1900 to 1904 and then to Oakland and the East Bay from 1907 to 1909. It has been present in the rodents of western North America ever since, as fear of the consequences of

1965-834: A pH of 7.2–7.6 (it still grows slowly), but can live in a large temperature and pH range. It dies very rapidly if exposed to a UV light, gets dried out, or in temperatures higher than 40°C. There are 11 species in the Yersinia genus, and three of them cause human diseases. The other two are Yersinia pseudotuberculosis and Yersinia enterocolitica , infections by either of these are usually acquired from ingesting contaminated food or water. Several complete genome sequences are available for various strains and subspecies of Y. pestis : strain KIM (of biovar Y. p. medievalis ), and strain CO92 (of biovar Y. p. orientalis , obtained from

2096-538: A policy of internal reconciliation and tried to reorganize royal administration. The dukes yielded half their estates for the maintenance of the king and his court in Pavia. On the foreign affairs side, Authari managed to thwart the dangerous alliance between the Byzantines and the Franks. Authari died in 591 and was succeeded by Agilulf , the duke of Turin , who also married Theodelinda in the same year. Agilulf successfully fought

2227-631: A port in the northern languages ; consequently, Skiæren-Heal is the port of the Skiæren, a name well adapted to the port of Stockholm , in the Upplandske Skiæren, and the country may be justly called Scorung or Skiærunga. The legendary king Sceafa of Scandza was an ancient Lombardic king in Anglo-Saxon legend . The Old English poem Widsith , in a listing of famous kings and their countries, has Sceafa [weold] Longbeardum, so naming Sceafa as ruler of

2358-485: Is a facultative anaerobic organism that can infect humans primarily via the Oriental rat flea ( Xenopsylla cheopis ), but also through airborne droplets for its pneumonic form. Yersinia pestis is a parasite of its host, the rat flea , which is also a parasite of rats, hence Y. pestis is a hyperparasite . Y. pestis was discovered in 1894 by Alexandre Yersin , a Swiss/French physician and bacteriologist from

2489-594: Is a protein tyrosine phosphatase that contributes to the ability of Y. pestis to evade immune system cells. In macrophages, YopH has been shown to dephosphorylate p130Cas , Fyb ( FYN binding protein) SKAP-HOM and Pyk , a tyrosine kinase homologous to FAK . YopH also binds the p85 subunit of phosphoinositide 3-kinase , the Gab1 , the Gab2 adapter proteins, and the Vav guanine nucleotide exchange factor . YopE functions as

2620-428: Is a critical component of virulence for Yersinia species. Many novel sRNAs were identified from Y. pestis grown in vitro and in the infected lungs of mice suggesting they play role in bacterial physiology or pathogenesis. Among them sR035 predicted to pair with SD region and transcription initiation site of a thermo-sensitive regulator ymoA, and sR084 predicted to pair with fur, ferric uptake regulator . In

2751-456: Is an acetyltransferase that binds to a conserved α-helix of MAPK kinases . YopJ acetylates MAPK kinases at serines and threonines that are normally phosphorylated during activation of the MAP kinase cascade . YopJ is activated in eukaryotic cells by interaction with target cell phytic acid (IP6). This disruption of host cell protein kinase activity causes apoptosis of macrophages, and this

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2882-514: Is better to maintain liberty by arms than to stain it by the payment of tribute." The Vandals prepared for war and consulted Godan (the god Odin ), who answered that he would give victory to those whom he would see first at sunrise. The Winnili were fewer in number and Gambara sought help from Frea (the goddess Frigg ), who advised that all Winnili women should tie their hair in front of their faces like beards and march in line with their husbands. At sunrise, Frea turned her husband's bed so that he

3013-403: Is famous. Among other things, these virulence factors are required for bacterial adhesion and injection of proteins into the host cell, invasion of bacteria in the host cell (via a type-III secretion system ), and acquisition and binding of iron harvested from red blood cells (by siderophores ). Y. pestis is thought to be descended from Y. pseudotuberculosis , DNA studies have found that

3144-400: Is generally regarded as the first historically recorded epidemic of Yersinia pestis . This conclusion is based on historical descriptions of the clinical manifestations of the disease and the detection of Y. pestis DNA from human remains at ancient grave sites dated to that period. Genetic studies of modern and ancient Yersinia pestis DNA suggest that the origin of the Justinian plague

3275-669: Is now northern Germany, near the Elbe river. They migrated south, and by the end of the fifth century, the Lombards had moved into the area roughly coinciding with modern Austria and Slovakia north of the Danube . Here they subdued the Heruls and later fought frequent wars with the Gepids . The Lombard king Audoin defeated the Gepid leader Thurisind in 551 or 552, and Audoin's successor Alboin eventually destroyed

3406-412: Is proposed to be important for the establishment of infection and for evasion of the host immune response. YopO is a protein kinase also known as Yersinia protein kinase A (YpkA). YopO is a potent inducer of human macrophage apoptosis. It has also been suggested that a bacteriophage  – Ypφ – may have been responsible for increasing the virulence of this organism. Depending on which form of

3537-498: Is related to both Yersinia enterocolitica and Yersinia pseudotuberculosis , the pathogen from which Y. pestis evolved and responsible for the Far East scarlet-like fever . It causes the disease plague , which caused the Plague of Justinian and the Black Death , the deadliest pandemic in recorded history. Plague takes three main forms: pneumonic , septicemic , and bubonic . It

3668-518: Is responsible for current plague outbreaks. On January 15, 2018, researchers at the University of Oslo and the University of Ferrara suggested that humans and their parasites (most likely fleas and lice at the time) were the biggest carriers of the plague. In 2010, researchers in Germany definitively established, using PCR evidence from samples obtained from Black Death victims, that Y. pestis

3799-401: Is the first and the best known outbreak of the first plague pandemic, which continued to recur until the middle of the 8th century. Some historians believe the first plague pandemic was one of the deadliest pandemics in history , resulting in the deaths of an estimated 15–100 million people during two centuries of recurrence, a death toll equivalent to 25–60% of Europe's population at the time of

3930-776: Is the seventh-century Origo Gentis Langobardorum ( Origin of the Lombard People ). The Origo Gentis Langobardorum tells the story of a small tribe called the Winnili dwelling in Northern Germany/Denmark zone (the Historia Langobardorum codicis Gothani writes that the Winnili first dwelt near a river called Vindilicus on the extreme boundary of Gaul ). The Winnili were split into three groups and one part left their native land to seek foreign fields. The reason for

4061-1001: The Afanasievo culture in Siberia, the Corded Ware culture in Estonia, the Sintashta culture in Russia, the Unetice culture in Poland, and the Andronovo culture in Siberia. In 2018, the emergence and spread of the pathogen during the Neolithic decline (as far back as 6,000 years ago) was published. A site in Sweden was the source of the DNA evidence and trade networks were proposed as

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4192-567: The Avars and then to the Franks. Grimoald managed to regain control over the duchies and deflected the late attempt of the Byzantine emperor Constans II to conquer southern Italy. He also defeated the Franks. At Grimoald's death in 671 Perctarit returned and promoted tolerance between Arians and Catholics, but he could not defeat the Arian party, led by Arachi, duke of Trento , who submitted only to his son,

4323-516: The Battle of Taginae . In approximately 560, Audoin was succeeded by his son Alboin , a young and energetic leader who defeated the neighboring Gepidae and made them his subjects; in 566, he married Rosamund , daughter of the Gepid king Cunimund . In the same year, he made a pact with Khagan Bayan . Next year the Lombards and the Avars destroyed the Gepid kingdom in the Lombard–Gepid War ,

4454-565: The Cochrane Collaboration found no studies of sufficient quality to make any statement on the efficacy of the vaccine. In 1894, two bacteriologists, Alexandre Yersin of Switzerland and Kitasato Shibasaburō of Japan, independently isolated in Hong Kong the bacterium responsible for the 1894 Hong Kong plague . Though both investigators reported their findings, a series of confusing and contradictory statements by Kitasato eventually led to

4585-517: The County of Sicily . In this period, the southern part of Italy still under Lombard domination was known to the Norse as Langbarðaland ('land of the Lombards'), as inscribed in the Norse runestones . Their legacy is also apparent in the name of the region of Lombardy in northern Italy. According to their traditions, the Lombards initially called themselves the Winnili . After a reported major victory against

4716-613: The Early Christian Church . Gradually, they adopted Roman or Romanized titles, names, and traditions, and partially converted to orthodoxy (in the seventh century), though not without a long series of religious and ethnic conflicts. By the time Paul the Deacon was writing, the Lombard language, dress and even hairstyles had nearly all disappeared in toto . The whole Lombard territory was divided into 36 duchies, whose leaders settled in

4847-799: The Elbe shortly after the beginning of the Christian era, next to the Chauci . Strabo states that the Lombards dwelt on both sides of the Elbe. He treats them as a branch of the Suebi , and states that: Now as for the tribe of the Suebi, it is the largest, for it extends from the Rhenus to the Albis; and a part of them even dwells on the far side of the Albis, as, for instance, the Hermondori and

4978-513: The Historia between 787 and 796 he was a Catholic monk and devoted Christian . He thought the pagan stories of his people "silly" and "laughable". Paul explained that the name "Langobard" came from the length of their beards. A modern theory suggests that the name "Langobard" comes from Langbarðr , a name of Odin . Priester states that when the Winnili changed their name to "Lombards", they also changed their old agricultural fertility cult to

5109-588: The Italian Peninsula between 568 and 774. The medieval Lombard historian Paul the Deacon wrote in the History of the Lombards (written between 787 and 796) that the Lombards descended from a small tribe called the Winnili, who dwelt in northern Germany before migrating to seek new lands. Earlier Roman-era historians wrote of the Lombards in the first century AD as being one of the Suebian peoples, also from what

5240-621: The Lower Elbe as Langobardic . The burial sites are crematorial and are usually dated from the sixth century BC through the third century AD, so a settlement breakoff seems unlikely. The lands of the lower Elbe fall into the zone of the Jastorf Culture and became Elbe-Germanic , differing from the lands between Rhine , Weser , and the North Sea . Archaeological finds show that the Lombards were an agricultural people. Tacitus also counted

5371-512: The Marcomannic Wars , 6,000 Lombards and Obii (sometimes thought to be Ubii ) crossed the Danube and invaded Pannonia . The two tribes were defeated, whereupon they ceased their invasion and sent Ballomar, King of the Marcomanni, as ambassador to Aelius Bassus , who was then administering Pannonia. Peace was made and the two tribes returned to their homes, which in the case of the Lombards

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5502-590: The Pasteur Institute , during an epidemic of the plague in Hong Kong . Yersin was a member of the Pasteur school of thought. Kitasato Shibasaburō , a Japanese bacteriologist who practised Koch's methodology , was also engaged at the time in finding the causative agent of the plague. However, Yersin actually linked plague with a bacillus , initially named Pasteurella pestis ; it was renamed Yersinia pestis in 1944. Between one thousand and two thousand cases of

5633-548: The Ravenna Cosmography , Mauringa was the land east of the Elbe. The crossing into Mauringa was very difficult. The Assipitti (possibly the Usipetes ) denied them passage through their lands and a fight was arranged for the strongest man of each tribe. The Lombard was victorious, passage was granted, and the Lombards reached Mauringa. The Lombards departed from Mauringa and reached Golanda. Scholar Ludwig Schmidt thinks this

5764-587: The Vandals in the first century, they changed their name to Lombards . The name Winnili is generally translated as 'the wolves', related to the Proto-Germanic root *wulfaz 'wolf'. The name Lombard was reportedly derived from the distinctively long beards of the Lombards. It is probably a compound of the Proto-Germanic elements * langaz (long) and * bardaz (beard). According to their own legends,

5895-508: The Vandals in the region of Carthage and the Ostrogoths ' kingdom in Italy . He had invested heavily in the construction of great churches, such as Hagia Sophia . As the empire tried to fund the projects, the plague caused tax revenues to decline through the massive number of deaths and the disruption of agriculture and trade. Justinian swiftly enacted new legislation to deal more efficiently with

6026-465: The innate immune system , as discussed below. In addition, some Y. pestis strains are capable of interfering with immune signaling (e.g., by preventing the release of some cytokines ). Y. pestis proliferates inside lymph nodes , where it is able to avoid destruction by cells of the immune system such as macrophages . The ability of Y. pestis to inhibit phagocytosis allows it to grow in lymph nodes and cause lymphadenopathy . YopH

6157-431: The Dukes ") during which the dukes did not elect any king, a period regarded as a time of violence and disorder. In 586, threatened by a Frankish invasion, the dukes elected Cleph's son, Authari , as king. In 589, he married Theodelinda , daughter of Garibald I of Bavaria , the Duke of Bavaria . The Catholic Theodelinda was a friend of Pope Gregory I and pushed for Christianization. In the meantime, Authari embarked on

6288-490: The Gepids in 567. The Lombards also settled in Pannonia (modern-day Hungary). Near Szólád , archaeologists have unearthed burial sites of Lombard men and women buried together as families, unusual among Germanic peoples at the time. Contemporary traces have also been discovered of Mediterranean Greeks and a possible migrant from France. Following Alboin's victory over the Gepids, he led his people into northeastern Italy , which had become severely depopulated and devastated after

6419-413: The Justinian plague originated in or near that region. However, there would appear to be no mention of bubonic plague in China until the year 610. The Byzantine historian Procopius first reported the epidemic in 541 from the port of Pelusium , near Suez in Egypt. Two other first hand reports of the plague's ravages were by the Syriac church historian John of Ephesus and Evagrius Scholasticus , who

6550-405: The Justinian plague strain. Additionally, a skeleton found in Tian Shan dating to around 180 AD and identified as an "early Hun" was found to contain DNA from Yersinia pestis closely related to the Tian Shan strain basal ancestor of the Justinian plague strain German samples. This finding suggests that the expansion of nomadic peoples who moved across the Eurasian steppe , such as the Xiongnu and

6681-441: The Langobardi; and at the present time these latter, at least, have, to the last man, been driven in flight out of their country into the land on the far side of the river. Consistent with this, Suetonius wrote that Roman general Nero Claudius Drusus defeated a large force of Germans and drove some "to the farther side of the Albis (Elbe)" river. The German archaeologist Willi Wegewitz defined several Iron Age burial sites at

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6812-624: The Lombard given name Ansegranus ("he with the beard of the gods") shows that the Lombards had this idea of their chief deity. The same Old Norse root Barth or Barði, meaning "beard", is shared with the Heaðobards mentioned in both Beowulf and in Widsith , where they conflict with the Danes . They were possibly a branch of the Langobards . Alternatively, some etymological sources suggest an Old High German root, barta, meaning "axe" (and related to English halberd), while Edward Gibbon puts forth an alternative suggestion which argues that: ...Börde (or Börd) still signifies "a fertile plain by

6943-436: The Lombards as a remote and aggressive Suebian tribe, listing them between the Semnones on the Elbe, and the Nerthus -worshipping tribes whose land of rivers and forest stretched to the sea. Writing in the late first century AD, he described the Langobardi in his Germania saying that "their scanty numbers are a distinction" because "surrounded by a host of most powerful tribes, they are safe, not by submitting, but by daring

7074-399: The Lombards occurred between AD 9 and 16, by the Roman court historian Velleius Paterculus , who accompanied a Roman expedition as prefect of the cavalry. Paterculus says that under Tiberius the "power of the Langobardi was broken, a race surpassing even the Germans in savagery". From the combined testimony of Strabo (AD 20) and Tacitus (AD 117), the Lombards dwelt near the mouth of

7205-468: The Lombards or Winnili lived originally in the Vinuiloth (Vinovilith) mentioned by Jordanes , in his masterpiece Getica , to the north of Uppsala , Sweden. Scoringa was near the province of Uppland , so just north of Östergötland . The footnote then explains the etymology of the name Scoringa: The shores of Uppland and Östergötland are covered with small rocks and rocky islands , which are called in German Schæren and in Swedish Skiaeren. Heal signifies

7336-443: The Lombards originated in Northern Germany/Denmark zone including modern-day Denmark. The Germanic origins of the Lombards is supported by genetic, anthropological, archaeological and earlier literary evidence. A legendary account of Lombard origins, history, and practices is the Historia Langobardorum ( History of the Lombards ) of Paul the Deacon , written in the eighth century. Paul's chief source for Lombard origins, however,

7467-443: The Lombards, who could retain their own laws. Rothari's son Rodoald succeeded him in 652, still very young, and was killed by his opponents. At the death of King Aripert I in 661, the kingdom was split between his children Perctarit , who set his capital in Milan, and Godepert , who reigned from Pavia ( Ticinum ). Perctarit was overthrown by Grimoald , son of Gisulf, duke of Friuli and Benevento since 647. Perctarit fled to

7598-399: The Lombards. From the second century onwards, many of the Germanic tribes recorded as active during the Principate started to unite into bigger tribal unions, such as the Franks , Alamanni , Bavarii , and Saxons . The Lombards are not mentioned at first, perhaps because they were not initially on the border of Rome, or perhaps because they were subjected to a larger tribal union, like

7729-445: The Lombards. Similarities between Langobardic and Gothic migration traditions have been noted among scholars. These early migration legends suggest that a major shifting of tribes occurred sometime between the first and second century BC, which would coincide with the time that the Teutoni and Cimbri left their homelands in Northern Germany and migrated through central Germany, eventually invading Roman Italy. The first mention of

7860-449: The Pope, who was supporting the dukes of Spoleto and Benevento against him, and entered Rome in 772, the first Lombard king to do so. But when Pope Hadrian I called for help from the powerful Frankish king Charlemagne , Desiderius was defeated at Susa and besieged in Pavia , while his son Adelchis was forced to open the gates of Verona to Frankish troops. Desiderius surrendered in 774, and Charlemagne, in an utterly novel decision, took

7991-463: The Saxons. It is, however, highly probable that, when the bulk of the Lombards migrated, a considerable part remained behind and afterwards became absorbed by the Saxon tribes in the Elbe region, while the emigrants alone retained the name of Lombards. However, the Historia Langobardorum codicis Gothani states that the Lombards were subjected by the Saxons around 300 but rose up against them under their first king, Agelmund, who ruled for 30 years. In

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8122-416: The above-mentioned Suebic territories, east of the Angrivarii on the Weser , and south of the Chauci on the coast, probably indicating a Lombard expansion from the Elbe to the Rhine. This double mention has been interpreted as an editorial error by Gudmund Schütte, in his analysis of Ptolemy. However, the Historia Langobardorum codicis Gothani also mentions Patespruna ( Paderborn ) in connection with

8253-422: The acceptance of Yersin as the primary discoverer of the organism. Yersin named it Pasteurella pestis in honor of the Pasteur Institute , where he worked. In 1967, it was moved to a new genus and renamed Yersinia pestis in his honor. Yersin also noted that rats were affected by plague not only during plague epidemics, but also often preceding such epidemics in humans and that plague was regarded by many locals as

8384-565: The allies halved the prize of war and the nomads settled in Transylvania . In the spring of 568, Alboin, now fearing the aggressive Avars, led the Lombard migration into Italy , which he planned for years. According to the History of the Lombards, "Then the Langobards, having left Pannonia , hastened to take possession of Italy with their wives and children and all their goods." The Avars have agreed to shelter them if they wish to come back. Various other peoples who either voluntarily joined or were subjects of King Alboin were also part of

8515-442: The bacteria to pass the skin barrier. Y. pestis expresses a plasmin activator that is an important virulence factor for pneumonic plague and that might degrade on blood clots to facilitate systematic invasion. Many of the bacteria's virulence factors are antiphagocytic in nature. Two important antiphagocytic antigens , named F1 (fraction 1) and V or LcrV , are both important for virulence . These antigens are produced by

8646-410: The bacterium at normal human body temperature. Furthermore, Y. pestis survives and produces F1 and V antigens while it is residing within white blood cells such as monocytes , but not in neutrophils . Natural or induced immunity is achieved by the production of specific opsonic antibodies against F1 and V antigens; antibodies against F1 and V induce phagocytosis by neutrophils. In addition,

8777-406: The cause of the decline, and that there were three outbreaks in Northern Europe between 5,200 years ago and 4,900 years ago, with the final outbreak caused by a strain of Yersinia pestis with reshuffled genes. Between 1970 and 2020, 496 cases were reported in the United States. Cases have been found predominantly in New Mexico, Arizona, Colorado, California, Oregon, and Nevada. In 2008, plague

8908-401: The city by infected rats on grain ships arriving from Egypt. To feed its citizens, the city and outlying communities imported large amounts of grain, mostly from Egypt. The rat population in Egypt thrived on feeding from the large granaries maintained by the government, and the fleas thrived as well. Procopius, in a passage closely modelled on Thucydides , recorded that at its peak the plague

9039-437: The deaths of up to a quarter of the human population of the Eastern Mediterranean . Frequent subsequent waves of the plague continued to strike throughout the 6th, 7th and 8th centuries, with the disease becoming more localized and less virulent. A revisionist view expressed by scholars such as Lee Mordechai and Merle Eisenberg argues that the mortality of the Justinian Plague was far lower than previously believed. They say that

9170-407: The disagreements between the Pope and Byzantium concerning the reverence of icons , he annexed the Exarchate of Ravenna and the duchy of Rome . He also helped the Frankish marshal Charles Martel drive back the Arabs . The Slavs were defeated in the Battle of Lavariano , when they tried to conquer the Friulian Plain in 720. Liutprand's successor Aistulf conquered Ravenna for the Lombards for

9301-463: The disease. In Yunnan , China, inhabitants would flee from their homes as soon as they saw dead rats, and on the island of Formosa ( Taiwan ), residents considered the handling of dead rats heightened the risks of developing plague. These observations led him to suspect that the flea might be an intermediary factor in the transmission of plague, since people acquired plague only if they were in contact with rats that had died less than 24 hours before. In

9432-545: The exodus was probably overpopulation . The departing people were led by Gambara and her sons Ybor and Aio and arrived in the lands of Scoringa , perhaps the Baltic coast or the Bardengau on the banks of the Elbe . Scoringa was ruled by the Vandals and their chieftains, the brothers Ambri and Assi, who granted the Winnili a choice between tribute or war. The Winnili were young and brave and refused to pay tribute, saying "It

9563-497: The expected disease dynamics have not been found in any rodent. Several species of rodents are known to have a variable resistance, which could lead to an asymptomatic carrier status. Evidence indicates fleas from other mammals have a role in human plague outbreaks. The lack of knowledge of the dynamics of plague in mammal species is also true among susceptible rodents such as the black-tailed prairie dog ( Cynomys ludovicianus ), in which plague can cause colony collapse, resulting in

9694-507: The far north-western edges of Europe. In 2016, the Y. pestis bacterium was identified from DNA in teeth found at a Crossrail building site in London . The human remains were found to be victims of the Great Plague of London , which lasted from 1665 to 1666. In 2021, researchers found a 5,000-year-old victim of Y. pestis , the world's oldest-known, in hunter-gatherer remains in

9825-629: The first Lombard duchy, which he entrusted to his nephew Gisulf . Soon Vicenza , Verona and Brescia fell into Germanic hands. In the summer of 569, the Lombards conquered the main Roman centre of northern Italy , Milan . The area was then recovering from the terrible Gothic Wars , and the small Byzantine army left for its defence could do almost nothing. Longinus, the Exarch sent to Italy by Emperor Justin II , could only defend coastal cities that could be supplied by

9956-422: The first outbreak. Research published in 2019 argued that the 200-year-long pandemic's death toll and social effects have been exaggerated, comparing it to the modern third plague pandemic (1855–1960s). Furthermore, historians seek to play down the effects of the late-antique bubonic plague, arguing that our eyewitness accounts of the disease are hysterical in tone and misleading in effect. The plague of Justinian

10087-591: The first time but had to relinquish it when he was subsequently defeated by the king of the Franks, Pippin III , who was called by the Pope. After the death of Aistulf, Ratchis attempted to become king of Lombardy, but he was deposed by Desiderius , duke of Tuscany , the last Lombard to rule as king. Desiderius managed to take Ravenna definitively, ending the Byzantine presence in northern Italy. He decided to reopen struggles against

10218-555: The first to describe this phenomenon). Transmission of Y. pestis occurs during the futile attempts of the flea to feed. Ingested blood is pumped into the esophagus, where it dislodges bacteria lodged in the proventriculus, which is regurgitated back into the host circulatory system. Pathogenesis due to Y. pestis infection of mammalian hosts is due to several factors, including an ability of these bacteria to suppress and avoid normal immune system responses such as phagocytosis and antibody production. Flea bites allow for

10349-400: The flea digestive tract, among them the hemin storage system and Yersinia murine toxin (Ymt). Although Ymt is highly toxic to rodents and was once thought to be produced to ensure reinfection of new hosts, it is essential for flea colonization and for the survival of Y. pestis in fleas. The hemin storage system plays an important role in the transmission of Y. pestis back to

10480-524: The general south–north spread of the disease from Egypt into the rest of the Mediterranean world. It also explains why Sassanid Persia saw a later development of the outbreak despite stronger trade links with Central Asia. After samples of DNA from Yersinia pestis were isolated from skeletons of Justinian plague victims in Germany, it was found that modern strains currently found in the Tian Shan mountain range system are most basal known in comparison with

10611-632: The genetic evidence, including the suggestion that the plague may have entered Western Eurasia via more than one route, and perhaps struck England before Constantinople. According to 2024 research, major plagues that significantly impacted the Roman Empire, such as the Antonine Plague , the Plague of Cyprian , and the Plague of Justinian, are strongly linked to periods of cooler and drier climate conditions, indicating that colder weather may have contributed to

10742-518: The glut of inheritance suits being brought as a result of victims dying intestate . The plague's long-term effects on European and Christian history were enormous. As the disease spread to port cities around the Mediterranean, the struggling Goths were reinvigorated and their conflict with Constantinople entered a new phase. The plague weakened the Byzantine Empire at a critical point, when Justinian's armies had nearly retaken all of Italy and

10873-471: The increasing power of the Franks. In response, the kings tried to centralize power over time, but they definitively lost control over Spoleto and Benevento in the attempt. In 572, Alboin was murdered in Verona in a plot led by his wife, Rosamund, who later fled to Ravenna . His successor, Cleph , was also assassinated, after a ruthless reign of 18 months. His death began an interregnum of years (the " Rule of

11004-450: The infection is not clearly identified, which complicates prevention and early-warning programs. One such example was seen in a 2003 outbreak in Algeria . The transmission of Y. pestis by fleas is well characterized. Initial acquisition of Y. pestis by the vector occurs during feeding on an infected animal. Several proteins then contribute to the maintenance of the bacteria in

11135-607: The land to the west of the Soča River, namely the Gorizia Hills and the Venetian Slovenia . A new ethnic border was established that has lasted for over 1200 years up until the present time. The Lombard reign began to recover only with Liutprand the Lombard (king from 712), son of Ansprand and successor of the brutal Aripert II . He managed to regain a certain control over Spoleto and Benevento, and, taking advantage of

11266-477: The later Huns , had a role in spreading plague to West Eurasia from an origin in Central Asia. Earlier samples of Yersinia pestis DNA have been found in skeletons dating from 3000 to 800 BC, across West and East Eurasia. The strain of Yersinia pestis responsible for the Black Death , the devastating pandemic of bubonic plague , does not appear to be a direct descendant of the Justinian plague strain. However,

11397-756: The likely avenue of spread rather than migrations of populations. There is evidence that suggests Y. pestis may have originated in Europe in the Cucuteni–Trypillia culture , not in Asia as is more commonly believed. DNA evidence published in 2015 indicates Y. pestis infected humans 5,000 years ago in Bronze Age Eurasia, but genetic changes that made it highly virulent did not occur until about 4,000 years ago. The highly virulent version capable of transmission by fleas through rodents, humans, and other mammals

11528-586: The long Gothic War (535–554) between the Byzantine Empire and the Ostrogothic Kingdom . The Lombards were joined by numerous Saxons , Heruls , Gepids, Bulgars , Thuringians and Ostrogoths , and their invasion of Italy was almost unopposed. By late 569, they had conquered all of northern Italy and the principal cities north of the Po River except Pavia , which fell in 572. At the same time, they occupied areas in central and southern Italy. They established

11659-413: The main cities. The king ruled over them and administered the land through emissaries called gastaldi . This subdivision, however, together with the independent indocility of the duchies, deprived the kingdom of unity, making it weak even when compared to the Byzantines, especially since these had begun to recover from the initial invasion. This weakness became even more evident when the Lombards had to face

11790-697: The market by the Food and Drug Administration . It was of limited effectiveness and could cause severe inflammation . Experiments with genetic engineering of a vaccine based on F1 and V antigens are underway and show promise. However, bacteria lacking antigen F1 are still virulent, and the V antigens are sufficiently variable such that vaccines composed of these antigens may not be fully protective. The United States Army Medical Research Institute of Infectious Diseases has found that an experimental F1/V antigen-based vaccine protects crab-eating macaques , but fails to protect African green monkey species . A systematic review by

11921-438: The migration. Whence, even until today, we call the villages in which they dwell Gepidan , Bulgarian , Sarmatian , Pannonian , Suabian , Norican , or by other names of this kind." At least 20,000 Saxon warriors, old allies of the Lombards, and their families joined them in their new migration. The first important city to fall was Forum Iulii ( Cividale del Friuli ) in northeastern Italy , in 569. There, Alboin created

12052-468: The modern Latvian and Estonian border area. Between 5,300 and 4,900 YBP the population of neolithic farmers in northern Europe underwent a marked decline. It had not been determined whether this was the result of agricultural recession or from Y. pestis infection within the population. A 2024 study of Neolithic graves in Denmark and western Sweden concluded that plague was sufficiently widespread to be

12183-515: The most energetic of all Lombard kings. He extended his dominions, conquering Liguria in 643 and the remaining part of the Byzantine territories of inner Veneto , including the Roman city of Opitergium ( Oderzo ). Rothari also made the famous edict bearing his name, the Edictum Rothari , which established the laws and the customs of his people in Latin : the edict did not apply to the tributaries of

12314-565: The old lands of the Burgundes . In Vurgundaib, the Lombards were stormed in camp by " Bulgars " (probably Huns ) and were defeated; King Agelmund was killed and Laimicho was made king. He was in his youth and desired to avenge the slaughter of Agelmund. The Lombards themselves were probably made subjects of the Huns after the defeat but rose up and defeated them with great slaughter, gaining great booty and confidence as they "became bolder in undertaking

12445-432: The other Yersinia species. pFra codes for a phospholipase D that is important for the ability of Y. pestis to be transmitted by fleas. pPla codes for a protease , Pla, that activates plasmin in human hosts and is a very important virulence factor for pneumonic plague. Together, these plasmids, and a pathogenicity island called HPI, encode several proteins that cause the pathogenesis for which Y. pestis

12576-565: The outbreak on trade caused authorities to hide the dead of the Chinatown residents long enough for the disease to be passed to widespread species of native rodents in outlying areas. Three main strains are recognised: Y. p. antiqua , which caused a plague pandemic in the sixth century; Y. p. medievalis , which caused the Black Death and subsequent epidemics during the second pandemic wave; and Y. p. orientalis , which

12707-404: The overall death toll is based on an estimate. In his Secret History , he records the devastation in the countryside and reports the ruthless response by the hard-pressed Justinian: When pestilence swept through the whole known world and notably the Roman Empire, wiping out most of the farming community and of necessity leaving a trail of desolation in its wake, Justinian showed no mercy towards

12838-513: The perils of war". Tacitus also noted that the Lombards were subjects of Marobod the King of the Marcomanni , who was allied with Rome when Arminius and his allies won the Battle of Teutoburg Forest in 9 AD. However, after the outbreak of war between Arminius and Marobod in 17 AD the Lombards and Semnones switched to the alliance of Arminius. They detested Marobod's title of king, and saw Arminius as

12969-716: The philo-Catholic Cunincpert . The Lombards engaged in fierce battles with Slavic peoples during these years: from 623 to 626 the Lombards unsuccessfully attacked the Carantanians , and, in 663–64, the Slavs raided the Vipava Valley and the Friuli . Religious strife and the Slavic raids remained a source of struggle in the following years. In 705, the Friuli Lombards were defeated and lost

13100-636: The plague are still reported to the World Health Organization every year. With proper antibiotic treatment, the prognosis for victims is much better than before antibiotics were developed. Cases in Asia increased five- to six-fold during the time of the Vietnam War , possibly due to the disruption of ecosystems and closer proximity between people and animals. The plague is now commonly found in sub-Saharan Africa and Madagascar, areas that now account for over 95% of reported cases. The plague also has

13231-507: The plague infects the individual, the plague develops a different illness; however, the plague overall affects the host cell's ability to communicate with the immune system, hindering the body bringing phagocytic cells to the area of infection. Y. pestis is a versatile killer. In addition to rodents and humans, it is known to have killed camels, chickens, and pigs. Domestic dogs and cats are susceptible to plague, as well, but cats are more likely to develop illness when infected. In either,

13362-441: The plague might have caused high mortality in specific places, but it did not cause widespread demographic decline or decimate Mediterranean populations. According to them, any direct mid-to-long term effects of plague were minor. However, their position has been the subject of a concerted critique by Peter Sarris, who challenged both their core methodology and their handling of the sources. Sarris also provides up-to-date discussion of

13493-527: The plague, and plague victims at an early Anglo-Saxon burial site at Edix Hill near Cambridge show that it also reached Britain. Procopius said that plague sufferers experienced delusions, nightmares, fevers, swellings in the groin, armpits and behind the ears, and coma or death. Treatments included cold baths, powders "blessed" by saints, magic amulets or rings, and various drugs, especially alkaloids. When these treatments failed, people went to hospitals or tried to quarantine themselves. The Plague of Justinian

13624-478: The pneumonic form, humans can spread the bacterium to others through airborne respiratory droplets; others who catch plague this way will mostly contract the pneumonic form themselves. Several species of rodents serve as the main reservoir for Y. pestis in the environment. In the steppes , the natural reservoir is believed to be principally the marmot . In the western United States, several species of rodents are thought to maintain Y. pestis . However,

13755-401: The powerful Byzantine fleet. Pavia fell after a siege of three years, in 572, becoming the first capital city of the new Lombard kingdom of Italy. In the following years, the Lombards penetrated further south, conquering Tuscany and establishing two duchies, Spoleto and Benevento under Zotto , which soon became semi-independent and even outlasted the northern kingdom, surviving well into

13886-428: The rebel dukes of northern Italy, conquering Padua in 601, Cremona and Mantua in 603, and forcing the Exarch of Ravenna to pay tribute. Agilulf died in 616; Theodelinda reigned alone until 628 when she was succeeded by Adaloald . Arioald , the head of the Arian opposition who had married Theodelinda's daughter Gundeperga, later deposed Adaloald. Arioald was succeeded by Rothari , regarded by many authorities as

14017-459: The ruined freeholders. Even then, he did not refrain from demanding the annual tax, not only the amount at which he assessed each individual, but also the amount for which his deceased neighbors were liable. As a result of the plague in the countryside, farmers could not take care of crops and the price of grain rose in Constantinople. Justinian had expended huge amounts of money for wars against

14148-399: The second half of the fourth century, the Lombards left their homes, probably due to bad harvests, and embarked on their migration. The migration route of the Lombards in 489, from their homeland to " Rugiland ", encompassed several places: Scoringa (believed to be their land on the Elbe shores), Mauringa , Golanda , Anthaib , Banthaib , and Vurgundaib ( Burgundaib ). According to

14279-501: The side of a river," and a district near Magdeburg is still called the lange Börde. According to this view Langobardi would signify "inhabitants of the long bord of the river;" and traces of their name are supposed still to occur in such names as Bardengau and Bardewick in the neighborhood of the Elbe. According to the Gallaecian Christian priest , historian and theologian Paulus Orosius (translated by Daines Barrington ),

14410-413: The spread of Justinian plague may have caused the evolutionary radiation that gave rise to the currently extant 0ANT.1 clade of strains. The number of deaths is uncertain. Some modern scholars believe that the plague killed up to 5,000 people per day in Constantinople at the peak of the pandemic. According to one view, the initial plague ultimately killed perhaps 40% of the city's inhabitants and caused

14541-432: The spread of these diseases during that time. It is thought climate stress interacted with social and biological variables, such as food availability, rodent populations, and human migration, making populations more susceptible to disease. Yersinia pestis Yersinia pestis ( Y. pestis ; formerly Pasteurella pestis ) is a gram-negative , non-motile , coccobacillus bacterium without spores that

14672-408: The symptoms are similar to those experienced by humans, and can be deadly to the animal. People can be exposed by coming into contact with an infected animal (dead or alive), or inhaling infectious droplets that a sick dog or cat has coughed into the air. A formalin -inactivated vaccine was available in the United States for adults in 1993 at high risk of contracting the plague until removal from

14803-593: The toils of war." During the reign of King Claffo , the Langobards occupied parts of modern-day Upper and Lower Austria and converted to Arian Christianity . In 505 the Herulians attacked and defeated them, obliging them to pay tax and withdraw to Northern Bohemia . In 508, King Rodulf sent his brother to the Lombard court to collect tribute and extend the truce; however, he was stabbed by Rometrud, sister of King Tato . Rodulf personally led his forces against Tato, but

14934-534: The twelfth century. Wherever they went, they were joined by the Ostrogothic population, which was allowed to live peacefully in Italy with their Rugian allies under Roman sovereignty. The Byzantines managed to retain control of the area of Ravenna and Rome, linked by a thin corridor running through Perugia . When they entered Italy, some Lombards retained their native form of paganism , while some were Arian Christians. Hence they did not enjoy good relations with

15065-521: The twelve Mongolian autonomous regions in Northern China. The patient received treatment, and 28 symptomless contacts were placed in quarantine. In July 2020, officials increased precautions after a case of bubonic plague was confirmed in Bayannur , a city in China's Inner Mongolia autonomous region. The patient was quarantined and treated. According to China's Global Times , a second suspected case

15196-552: The two are 83% similar, which is high enough to be considered the same species. In 1981 it was proposed that Y. pestis be reclassified as a subspecies of Y. pseudotuberculosis , but the Judicial Commission of the International Committee on Systematic Bacteriology declined to do this because the course of Y. pestis disease is so different than that of Y. pseudotuberculosis , which usually causes

15327-516: The type-III secretion system (T3SS) allows Y. pestis to inject proteins into macrophages and other immune cells. These T3SS-injected proteins, called Yersinia outer proteins (Yops), include Yop B/D, which form pores in the host cell membrane and have been linked to cytolysis . The YopO, YopH , YopM, YopT, YopJ, and YopE are injected into the cytoplasm of host cells by T3SS into the pore created in part by YopB and YopD. The injected Yops limit phagocytosis and cell signaling pathways important in

15458-428: The urban and sylvatic (forest) cycles of Y. pestis, most of the spreading occurs between rodents and fleas. In the sylvatic cycle, the rodent is wild, but in the urban cycle, the rodent is primarily the brown rat ( Rattus norvegicus ). In addition, Y. pestis can spread from the urban environment and back. Transmission to humans is usually through the bite of infected fleas. If the disease has progressed to

15589-448: The virulence genes. Some 63 novel putative sRNAs were identified through deep sequencing of the Y. pestis sRNA-ome. Among them was Yersinia -specific (also present in Y. pseudotuberculosis and Y. enterocolitica ) Ysr141 ( Yersinia small RNA 141). Ysr141 sRNA was shown to regulate the synthesis of the type III secretion system (T3SS) effector protein YopJ. The Yop-Ysc T3SS

15720-647: The western Mediterranean coast; the evolving conquest would have reunited the core of the Western Roman Empire with the Eastern Roman Empire. Although the conquest occurred in 554, the reunification did not last long. In 568, the Lombards invaded Northern Italy , defeated the small Byzantine army that had been left behind and established the Kingdom of the Lombards . Gaul is known to have suffered severely from

15851-535: Was a child in Antioch at the time and later became a church historian. Evagrius was afflicted with the buboes associated with the disease but survived. During the disease's four returns in his lifetime, he lost his wife, a daughter and her child, other children, most of his servants and people from his country estate. According to contemporary sources, the outbreak in Constantinople was thought to have been carried to

15982-408: Was also investigated, and a level 3 alert was issued, in effect until the end of the year. It forbade hunting and eating of animals that could carry plague, and called on the public to report suspected cases. Lombards The Lombards ( / ˈ l ɒ m b ər d z , - b ɑːr d z , ˈ l ʌ m -/ ) or Longobards ( Latin : Longobardi ) were a Germanic people who conquered most of

16113-466: Was ambushed and killed from a hill. In the 540s, Audoin (ruled 546–560) led the Lombards across the Danube once more into Pannonia . Thurisind , King of the Gepids attempted to expel them, and both peoples asked for help from the Byzantines. Justinian I sent his army against the Gepids; however, it was routed on the way by the Herulians and the sides signed a two-year truce. Revenging what he felt as

16244-566: Was commonly found in sub-Saharan Africa and Madagascar, areas that accounted for over 95% of the reported cases. In September 2009, the death of Malcolm Casadaban , a molecular genetics professor at the University of Chicago , was linked to his work on a weakened laboratory strain of Y. pestis . Hemochromatosis was hypothesised to be a predisposing factor in Casadaban's death from this attenuated strain used for research. On November 3, 2019, two cases of pneumonic plague were diagnosed at

16375-453: Was facing east, and woke him. So Godan spotted the Winnili first and asked, "Who are these long-beards?," and Frea replied, "My lord, thou hast given them the name, now give them also the victory." From that moment onwards, the Winnili were known as the Longbeards (Latinised as Langobardi , Italianised as Longobardi , and Anglicized as Langobards or Lombards ). When Paul the Deacon wrote

16506-907: Was found in two individuals associated with the Srubnaya culture from the Samara region in Russia from around 3,800 years ago and an Iron Age individual from Kapan , Armenia, from around 2,900 years ago. This indicates that at least two lineages of Y. pestis were circulating during the Bronze Age in Eurasia. The Y. pestis bacterium has a relatively large number of nonfunctioning genes and three "ungainly" plasmids, suggesting an origin less than 20,000 years ago. One such strain has been identified from about 4000  BP (the "LNBA lineage" (Late Neolithic and Bronze Age lineage)) in western Britain, indicating that this highly transmissible form spread from Eurasia to

16637-599: Was further east, perhaps on the right bank of the Oder . Schmidt considers the name the equivalent of Gotland , meaning simply "good land". This theory is highly plausible; Paul the Deacon mentions the Lombards crossing a river, and they could have reached Rugiland from the Upper Oder area via the Moravian Gate . Moving out of Golanda, the Lombards passed through Anthaib and Banthaib until they reached Vurgundaib, believed to be

16768-588: Was in Central Asia . The most basal or root level existing strains of the Yersinia pestis as a whole species are found in Qinghai , China. Other scholars contest that, rather than Central Asia, the specific strain that composed the Justinian plague began in sub-Saharan Africa, and that the plague was spread to the Mediterranean by merchants from the Kingdom of Aksum in East Africa. This point of origin aligns more with

16899-474: Was killing 10,000 people in Constantinople daily, but the accuracy of the figure is in question, and the true number will probably never be known. He noted that because there was no room to bury the dead, bodies were left stacked in the open. Funeral rites were often left unattended to, and the entire city smelled like the dead. Given such circumstances, it is highly probable that a sudden increase in mortality rates may not have been as accurately recorded, hence why

17030-430: Was the cause of the medieval Black Death . In 2011, the first genome of Y. pestis isolated from Black Death victims was published, and concluded that this medieval strain was ancestral to most modern forms of Y. pestis . In 2015, Cell published results from a study of ancient graves. Plasmids of Y. pestis were detected in archaeological samples of the teeth of seven Bronze Age individuals, in

17161-667: Was the lands of the lower Elbe. In the mid-second century, the Lombards supposedly appeared in the Rhineland , because according to Claudius Ptolemy , the Suebic Lombards lived "below" the Bructeri and Sugambri , and between these and the Tencteri . To their east stretching northwards to the central Elbe are the Suebi Angili . But Ptolemy also mentions the "Laccobardi" to the north of

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